Novitates Gabonenses 94 Salpingifera (Achariaceae) a new monotypic and endemic genus from Gabon

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Blumea 69, 2024: 196–200 www.ingentaconnect.com/content/nhn/blumea https://doi.org/10.3767/blumea.2024.69.02.07 RESEARCH ARTICLE Citation:BretelerFJ,WieringaJJ.2024.NovitatesGabonenses94.Salpingifera(Achariaceae)anewmonotypicandendemicgenusfromGabon. Blumea69(2):196–200.https://doi.org/10.3767/blumea.2024.69.02.07. Effectivelypublishedonline:30October2024. INTRODUCTION In 1923 Pellegrin described a new species of Lindackeria C.Presl(Flacourtiaceae)fromAfricaandnameditL. ngounyensisPellegr.aftertheNgounyeRiverbasininCentralGabon. ThetypematerialcollectedbyLeTestuisonlyflowering. WhenSleumer(1974a)revisedthegenusCamptostylusGilg (Flacourtiaceae, now Achariaceae)hestatedthatLindackeria ngounyensisprobably(‘vermutlich’)belongedtothisgenus.The only argument was that the leaves resembled those of C. ovalis Chipp, but because only staminate flowers were known he furtherignoredthespeciesinhisrevision.Maybethepresenceof staminate,sounisexual,flowerswasanargumentforSleumer aswell,butsuchisnotstatedexplicitly.Inhisfollowingrevision of the African Lindackeriaspecies(Sleumer1974b)heexcluded L. ngounyensis from this genus and, although fruits were still unknown, he strengthened his earlier statements now saying thatitalmostsurely(‘wohlsicher’)belongedtoCamptostylus. Without any argument Hul (1995) transferred all species of Camptostylus and LindackeriaoccurringinGabontoOncoba Forssk.andsoL. ngounyensis became Oncoba ngounyensis (Pellegr.)Hul.Ina later publication Hul & Breteler (1997) argumentedthattheprevioussegregationofgenerabyGilg (1925),thatwasfollowedbySleumer(1974b),wasbasedon mostly vague delimitations with a major dependency on fruit characters.Theymoreorlessrevertedbacktotheclassification byOliver(1868)intoawidelycircumscribedOncoba and transferredthenon-Gabonesespeciesasfarasnecessary. Chaseetal.(2002)divided theFlacourtiaceae genera over Salicaceae and Achariaceae. Oncoba s.str. was placed in Salicaceae, but the remaining genera of the Flacourtiaceae tribe Oncobeae, including those that were included in Oncoba byHul(1995),wenttoAchariaceae and were placed in a new tribe LindackerieaeZmarzty,undertheirformerpre-Hul(Hul 1995)genericnames,althoughnonewspecificcombinations weremade.Lindackerieae is predominantly an African tribe, currently including eight wholly African genera: Buchnerodendron Gürke, Caloncoba Gilg, Camptostylus, Grandidiera Jaub., Peterodendron Sleumer, Poggea Gürke, Prockiopsis Baill.and Xylotheca Hochst. Prockiopsis occurs only in Madagascar. ThreefurthergeneraareconfinedtoCentralandSouthAmerica: Carpotroche Endl.,MaynaAubl.andKuhlmanniodendron Fiaschi&Groppo(Groppoetal.2016).FinallyLindackeria as currently circumscribed occurs in both Africa and the Americas, theonlygenusofthetribethatdoesso.Thisclassificationis likelywrongandwillbethesubjectofanotherpaper(Breteler &Wieringainprep.). MATERIAL AND METHODS Based on herbarium and alcohol collections at BM, BR, BRLU,K,L,P,UandWAG(acronymsaccordingtoThiers continuouslyupdated),theleaves,flowers,fruitsandseedsof all Lindackerieaegenerahavebeenmorphologicallystudied. OftheAfricangeneraallspecieshavebeeninvestigated.The American genera have all been investigated, but the different speciesonlyasfarasmaterialwasavailableinL,UorWAG. RESULTS SinceLindackeria ngounyensis was included in the LindackerieaewhentheseweredescribedbyChaseetal.(2002),the speciesfitsthetribaldescription.Toincludethisspeciesthe description had to be stretched to accommodate for 4 sepals perflower.Thetribalcharacter‘inflorescencenotracemosespicate’isonlytruewhentheindividualflowersareseenas one-floweredcymes(seebelow).Thespeciesdoesnotfitany oftheothertribalcircumscriptionsbyChaseetal.(2002)bysevNovitates Gabonenses 94 Salpingifera (Achariaceae) a new monotypic and endemic genus from Gabon F.J.Breteler1,2,J.J.Wieringa2 1 FormerlyHerbariumVadense,Grintweg303,6708ARWageningen,The Netherlands. 2 NaturalisBiodiversityCenter,Darwinweg2,2333CRLeiden,TheNetherlands; corresponding author e-mail: [email protected]. Key words Achariaceae new genus taxonomy tropical Africa Abstract The aberrant Oncoba ngounyensisfromGabon,originallydescribedinLindackeria(thenFlacourtiaceae, nowadays Achariaceae),butexcludedfromthatgenusbySleumer,iscloserinvestigatedtoseewhetheritbelongs in Camptostylus as suggested by that author or in any other genus of the Lindackerieae.Itisconcludedthatthis species belongs to neither Lindackeria nor Camptostylus.WithintheLindackerieae, Oncoba ngounyensis occupies a morphologically isolated position which is taxonomically best expressed by placing it in a new genus of its own, we name Salpingifera. 197 F.J.Breteler&J.J.Wieringa:Salpingifera,newgenusfromGabon eral characters, except for Erythrospermeae.However,itonly fitstherebecausetheexceptions,invokedby‘often’or‘usually’ phrases,havetobechosenseveraltimestofitL.ngounyensis. Inthattribe‘thesepalsandpetalsoftenindistinguishablefrom oneanother’and‘petalsusuallywithanadnateadaxialbasal scale’donotnicelyaccommodatethisspecies with petals withoutanyscalesclearlydifferentfromthesepals.Wehave considered the genera included in Erythrospermeae based on genericdescriptionsandconcludedourspeciesdoesnotfit anyofthem. An important new character to separate groups within Lindackerieae is the presence of a resinous layer on young shoots, which usuallyispresentininflorescencesaswell.Thepresenceof such a resinous layer might hamper the development of stipules, bracts and bracteoles, so the linkage of the resinous layer with poorly or not developed stipules might rather be a functional correlationthanaclearphylogeneticsignal.Thisresinouslayer is present in the genera Caloncoba, Camptostylus, Xylotheca and in the American species of Lindackeria. Otherimportant characters are the number of sepals, the dehiscence of fruits, the presence of a sarcotesta and the flower arrangement within theinflorescence. The inflorescence of Lindackeria/Oncoba ngounyensis is different from that of the other genera of the Lindackerieae.The flowersseemtobearrangedinaraceme.Eachflowerisat base, on both sides of its pedicel, accompanied by a small, variablenumberoftrumpet-shapedorgans(Fig.3a,b).Pellegrin (1923)observedthatthesetrumpet-shapedorganslooklike pedicels of which the flowers are aborted or have not developedatall.However,asisevidentonphotostakeninthefield (Fig.3b),theseorgansareterminatedbyaglandproducinga liquidthatissearchedafterbyants.So,eitheritarepedicels of which each flower is extremely reduced to a gland, or it isacompletelyuniqueorganbearingglands.Interestingly, Lindacke rieae are supposed to have no disk glands, so it is peculiarthattheonlyremainoftheflowerwouldbeagland.Maybe some Lindackerieae do have some nectar production, which has never been observed, just like the glandular nature of these trumpet-shapedorganswasneverrecorded.Weconsidered these trumpet-shaped structures as pedicels with reduced flowers, which together with the single developed flower, are part of a dichasium of which only the top flower of the primary axishas fully been developed.Thatthesetrumpet-shaped organs are in fact reduced flowers in a dichasium is corroborated by the fact that they develop after the development of the floralbudandtheybeartheirownbractsandbracteoles.The wholeinflorescenceshouldbeconsideredasathyrse. Sleumer(1974a)consideredCamptostylus to have indehiscent fruits.However,asisevidentinsomeherbariumspecimens thesefruitsaretardilydehiscent.SinceCamptostylus has seeds with a fleshy seed coat this would be logical to expose these seeds.Lindackeria ngounyensis possesses the same fleshy seeds, but the fruits seem to be indehiscent, which is quite extraordinary for Lindackerieae.Indehiscentfruitswithinthis tribe have been described for Mayna and Carpotroche byGilg (1925),butasisevidentfromsomecarpologicalcollectionsin UofCarpotroche brasiliensis (Raddi)A.Gray(Ducke 3891), C. platyptera Pittier(Chatrou 86)andC. surinamensisUittien (Ek 796),atleastthefruitsofthesespeciesofCarpotroche are dehiscent,ormaybetardilyso.Possiblysomeotherspeciesof Carpotrocheareindeedindehiscent.Groppoetal.(2013)states for Carpotroche ‘maybetardilydehiscent’,whichisconfirmed byourobservations.Asfaraswecanassessfromherbarium collections Mayna fruitsareindeedindehiscent. Kulmanniodendron fruits are also described as indehiscent, but we have too poormaterialtoassessthisisindeedthecase.So,nexttoin L. ngounyensis, indehiscent fruits only occur in Mayna, likely Kulmanniodendron and maybe Carpotroche. Fleshy seed coats are present in Caloncoba, Camptostylus and likely in Carpotrochewhereherbariumlabelsmention‘arils’and Endlicher(1839:918)describesitas“Semina…epidermide carnosa”.Thesegenerahavedehiscentfruits,exceptthatwe cannot rule out that of some Carpotroche species the fruits may beindehiscent.Thisrendersthecombinationofindehiscent fruits with fleshy seed coats unique, only possibly matched in Carpotroche.FruitsoftheSouthAmericanCarpotroche can be easily distinguished from L. ngounyensisbythefree4–8styles wherethisAfricanspeciesonlyhasasinglestyle.Following theclassificationoffruittypesbySpjut(1994)L. ngounyensis fruitsshouldpossiblybeclassifiedasacarcerulus,although this fruit type does not specify whether the seeds should have asarcotestaornot.Thesefruitsseemsimilartotheindehiscent fruits of PunicaL.,whichalsodohaveasarcotestaandare classifiedasabalausta,butabalaustaincludesotherflower tissueslikeahypanthium,whichisnotthecasehere.Aspecific name for an indehiscent solely carpel-based fruit with seeds includingasarcotestadoesnotseemtoexist. Sleumer’s(1974a)statementthatLindackeria ngounyensis has to be excluded from (African) Lindackeria is approved. His sug gestion, however, that it should be placed in Camptostylus does not find any support in the morphological characters presentedinTable1.Theonlyactualmatchisthefleshyseed coat,acharacterthatwasnotknowntoSleumer. Lindackeria/Oncoba ngounyensis shows a unique combination of characters and cannot be placed in any of the existing genera of the Lindackerieae without distorting the current morphological genericcircumscriptions.ItistheonlyLindackeriae with four sepals.Moreover,theuniquetrumpet-shapedgland-likestructures ontheaxisoftheinflorescencestandoutasextraordinary.Also the inflorescence type – a reduced thyrse – is unique within the tribe.Theindehiscentfruit includingseedswithfleshy seed coatsis probablyauniquecharacterforthetribe.Thebest Camptostylus Lindackeria ngounyensis remaining African Lindackeria species Branchlets resinous not resinous not resinous Stipules small,resinous,obsoleteorabsent well-developed well-developed Leaf blades entire dentate dentate Inflorescence an elongate axillary axis with scattered, axillary, not resinous, with flowers in much axillary, paniculate, racemose, sometimes umbellate short, compact, resinous, few-flowered reduced thyrses or subumbellate; not resinous lateral racemes Sepals three four three Fruit tardily dehiscent, ± smooth or ribbed, indehiscent, smooth, many-seeded dehiscent, sometimes tardily so, spiny or bristly or many-seeded with short emergencies, few-to many-seeded Seed seedcoatcompletelyfleshy seedcoatcompletelyfleshy seedwiththickenedraphe,sometimesaslargeas the seed, developed into a basal aril or not Table 1MorphologicalcomparisonbetweenCamptostylus, Lindackeria ngounyensis, and the remaining African Lindackeriaspecies. 198 Blumea – Volume 69 / 2, 2024 solutionforthisspeciesistoplaceitinitsowngenus.This segregation is supported by unpublished molecular analyses (Zmarztyetal.pers.comm.).Weproposetoclassifyitinthe new monotypic genus Salpingifera: Salpingifera ngounyensis. CONCLUSION Lindackeria ngounyensis has to be excluded from African Lindackeria and cannot be placed in any other current genus of the Lindackeriae.Thisspeciesisbestclassifiedinaseparate, new genus named Salpingifera.Atpresent itisbestplaced in Lindackerieae, where the characters of this species have alreadybeenincludedinthedescriptionanddelimitation.Still, itsmorphologicaldescriptionmoreorlessfitsthatofErythrospermeaeaswell.AclassificationinErythrospermeae or even in a separate tribe could be considered when molecular data formoregenerabecomesavailableandtribalclassificationof someothergeneramayhavetobereconsidered. Fig. 1 Salpingifera ngounyensis(Pellegr.)Breteler&Wieringa.1.Floweringbranch;2.inflorescence;3.detailsofthetrumpet-shapedorgansontherachis; 4.flowerbud;5.sepal;6.petal;7.stamensandpistilode;8.pistillode;9.fruits;10.trumpet-shapedorgansatbaseoffruitingpedicel;11.seed(1–8:Le Testu 2235; 9–11: Breteler 10874).—DrawnbyG.Chypre,reproducedfromHul(1995). 199 F.J.Breteler&J.J.Wieringa:Salpingifera,newgenusfromGabon TAXONOMY SalpingiferaBreteler&Wieringa,gen. nov. Etymology.Thisnew, monotypic genus isnamedafter the unique trumpet-shapedorgansintheinflorescence.ItisderivedfromtheGreek wordσαλπιγξ(salpinx)havingsalping-asgenitivestem.SoSalpingifera meanstrumpet-bearing.Salpingiferahasthefemalegender. Type: Lindackeria ngounyensisPellegr. Shrub.Leaves alternate, stipulate, penninerved, margin dentate.Inflorescence axillary, racemose compound of dichasia (thyrse)reducedtoasingleflowerflankedbypedicelsbearing trumpet-shapedglands.Flowers actinomorphic, staminate or bisexual/pistillate? Sepals4,free,imbricate.Petals8–10,free, imbricate.Stamens free, many, with long anthers opening by 2lengthwiseslits.Ovarysuperior,unilocular,with5parietal, multiovulateplacentas.Fruit smooth, leathery, indehiscent, acarcerulus.Seedsmany,withcompletefleshyseedcoat. Embryosmall,embeddedincopiousendosperm.  Distribution—Monotypic,endemictoGabon. Salpingifera ngounyensis(Pellegr.) Breteler & Wieringa, comb. nov.—Fig.1–4;Map1 Lindackeria ngounyensisPellegr.,BulletinduMuséumNationald’Histoire Naturelle de Paris29 (1923) 591.—Oncoba ngounyensis(Pellegr.) Hul(1995)150.—Type: Le Testu 2235(holoP;isoBR,WAG),Gabon, Ngounié,16Oct.1917. Shrubs,2–3(–5) m tall. Indumentum: branchlets, stipules, petioles and inflorescences puberulous to shortly velutinous, denselysoor not.Stipules subulate, 1.5–4(–5)mmlong, caducous.Leaves: petiole (sub)terete, swollen and usually curvedatapex,0.5–5(–8.5)cmlong;laminaobovate,rarely elliptic,(11–)15–25(–33)by(4.5–)6–12(–15)cm,2–3times as long as wide, papery to coriaceous, margin dentate mainly in upper half and often obscurely so, base cuneate to narrowly roundedtonarrowlysubcordate, apex (0.5–)1–2.5cm long acuminate; glabrous above, glabrous to sparsely puberulous beneath;midribandthe9–12(–13)pairsofmainlateralnerves slightlyprominentabove,distinctlysobeneath.Inflorescence erect,upto8cmlong,fewtomanyflowered;pedunclevery short or absent; each dichasium with a single flower and laterally c.3mmlongglandsonpedicels.Bractstriangular,0.5–1mm long,hairy;bracteolesnarrowlytriangular,c.0.4mmlong,hairy. Flowersstaminateorbisexual/pistillate?(seenotes).Staminate flowers:pedicelfirm, articulatedatthebase, 3–5mmlong, puberulous;sepals4,free,oblong-elliptic,7–8by3–3.5mm, decussate, imbricate, puberulous outside, glabrous inside; petals6–8,narrowlyelliptictooblong,8–10by2–3mm,imbricate,free,glabrous,white;stamens25–30,free,anthers yellow,c.3mmlong,filamentswhite,2–2.5mmlong;pistillode absentorminute,c.2mmlong,glabrous.Pistillate or bisexual flowersunknown.Young fruitwithastyleofatleast4mm.Fruit ellipsoid,beaked,3–5by1.5–4cm,glabrous,sometimesvery slightly angled lengthwise, yellow to orange at maturity; style basepersistent,developedintoamucron.  Distribution—OnlyknownfromGabon.  Habitat&Ecology—Primaryorsecondaryevergreenforest, in dry land forest as well as in periodically inundated forest, at sea-leveltoc.100maltitude. Other specimens examined.Gabon,Moyen-Ogooué,Bidault et al.941 (WAG),Mabounié,45kmSEofLambaréné,nearNgouniéR.,18Oct.2012; Bidault et al.986(WAG),ibid.,25Oct.2012;Bidault et al. 1020(WAG), ibid.,1Feb.2013;Bidault et al. 1057(WAG),ibid.,5Feb2013;Bidault et al. 1290(BR,WAG),ibid.,14Nov.2013;Breteler 10874(P,WAG),Mbéné, c.30kmonBifoun–LambarénéRd.,3Mar.1992;Lachenaud et al. 1598 (BR,WAG),nearEzangaLake,24Nov.2013;Lachenaud et al. 2032(BR), nearOnanguéLake,23Oct.2014;McPherson 15232(BR,WAG),near EzangaLake,9Feb.1991;Sonké et al. 6068(BR),sin.loc.,16Oct.2012; van Nek 589(WAG),c.20kmSofEzanga,31Jan.1991;Wilks 2445(WAG) Ezanga,1991.  Notes—Judgedbyseveralfieldnotesthatdescribeboth staminateflowersand(young)fruits,itseemsstaminateand pistillate or bisexual flowers may be present on the same individual.However,allthesheetswesawbeareitherstaminate flowersorfruits.Sinceitisnot100%surethefieldnotesare based on single individuals we cannot be sure the species is monoeciousordioecious. ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! (! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( ! ( Map. 1 Distribution of Salpingifera ngounyensis (Pellegr.) Breteler & Wieringa. Fig. 2 Salpingifera ngounyensis (Pellegr.)Breteler&Wieringa.Olderstaminateflower(Bidault 941).—PhotobyE.Bidault. 200 Blumea – Volume 69 / 2, 2024 GroppoM,FavarettoBSG,DaSilvaCI,etal.2013.AnewspeciesofKuhlmanniodendron(Lindackerieae,Achariaceae)fromEasternBrazilandthe systematicpositionofthegenusinAchariaceae.SystematicBotany38: 162–171.https://doi.org/10.1600/036364413X662114. GroppoM,FiaschiP,SalatinoMLF,etal.2016.PlacementofKuhlmanniodendronFiaschi&GroppoinLindackeriae (Achariaceae,Malpighiales) confirmedbyanalysesofrbcLsequences,withnotesonpollenmorphology andwoodanatomy.PlantSystematicsandEvolution286:27–37.https:// doi.org/10.1007/s00606-010-0276-3. HulS.1995.FlacourtiaceaeIn:MoratP(ed),FloreduGabon34:1–82. MuséumNationald’HistoireNaturelle,Paris HulS,BretelerFJ.1997.RéductionsgénériquesdanslesOncobeae(Flacourtiaceae).Adansonia19:253–262. OliverD.1868.Bixineae.FloraofTropicalAfrica1:112–123.CoventGarden,London. PellegrinF.1923.PlantaeLetestuanaenovaeouPlantesnouvellesrécoltées parM.LeTestude1907à1919dansleMayombecongolaisVIII.Bulletin duMuséumNationald’HistoireNaturelledeParis29:591–593. SleumerH.1974a.RevisionderGattungCamptostylusGilg(Flacourtiaceae). BotanischeJahrbücherfürSystematiek,PflanzengeschichteundPflanzengeographie94(3):283–288. SleumerH. 1974b. DieafrikanischenArtenderGattung Lindackeria Presl(Flacourtiaceae).BotanischeJahrbücherfürSystematiek,PflanzengeschichteundPflanzengeographie94(3):311–326. SpjutRW.1994.Asystematictreatmentoffruittypes.MemoirsoftheNew YorkBotanicalGarden70:1–182. Thiers BM. Continuously updated. Index Herbariorum. https://sweetgum. nybg.org/science/ih. Fig. 3 Salpingifera ngounyensis (Pellegr.)Breteler &Wieringa.a.Inflorescencewithbuds;b.detailof Fig.2,showingtrumpet-likeglandularorgansinthe inflorescencewith ants drinkingexudate(Bidault 941).—PhotosbyE.Bidault. b a b a Fig. 4 Salpingifera ngounyensis (Pellegr.)Breteler&Wieringa.a.Inflorescencewithbudsandfreshmaleflowers;b.branchwithfruits(a:Bidault 441; b: Bidault 1584).—PhotosbyE.Bidault. Wehaveonlystudiedstaminateflowers.Thereforewedonot know whether fruits develop from bisexual or from pistillate flowers.Likely,whenthespeciesisdioecious,pistillateplants flower less abundantly and/or less frequently than staminate plantsandarelesslikelytobecollectedbybotanists. AcknowledgementsTheauthorsaregratefulto the Muséum National d’HistoireNaturelleinParisforpermissiontousetheillustrationbyG.Chypre of Oncoba ngounyensispublishedintheFloreduGabon.TheykindlyacknowledgealsothepermissionbyE.Bidaulttousehisexcellentandmost usefulfieldphotosandAglaiaBoumafordigitalimprovementoftwo.Thanks gotoSueZmarztywho,byhercomments,improvedthemanuscript.Sheis also acknowledged for her fruitful discussions about the generic delimitation in the Lindackerieae.FrankBroekeassistedinthecorrectusuageofthe Greekwordsalpinx.ThanksgoalsotoThomasZandvlietwhopreparedthe firstelectronicversionofthemanuscript. REFERENCES ChaseMW,ZmarztyS,DoloresLledoM,etal.2002.Whenindoubt,putit in Flacourtiaceae: a molecular phylogenetic analysis based on plastid rbcL DNAsequences.KewBulletin57(1):141–181. 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